Are the Effects of Use and Disuse Inherited? Part 4

[36] _Variation of Animals and Plants under Domestication_, i. 183, 186.

[37] _Variation of Animals and Plants under Domestication_, i. 130, 135; ii. 288.

[38] _Encyclopaedia Britannica_, article "Zoology."

[39] _Variation of Animals and Plants under Domestication_, ii. 367.

[40] _Variation of Animals and Plants under Domestication_, ii. 367. Why then does the cheetah inherit ancestral habits so inadequately that it is useless for the chase unless it has first learned to hunt for itself before being captured? (ii. 133).

[41] _Descent of Man_, p. 33.

[42] _Origin of Species_, pp. 210, 211.

[43] E. S. Delamer on _Pigeons and Rabbits_, pp. 132, 103. For other points referred to, see pages 133, 102, 100, 95, 131.

[44] _Origin of Species_, pp. 188, 110; _Descent of Man_, pp. 32-35; _Variation of Animals and Plants under Domestication_, ii. 289, 293. Use or disuse during lifetime of course co-operates, and in some cases, as in that of the canoe Indians, may be the princ.i.p.al or even perhaps the _sole_ cause of the change.

[45] For the importance of panmixia as invalidating Darwin's strongest evidence for use-inheritance--namely, that drawn from the effects of disuse in highly-fed domestic animals where there is supposed to be no economy of growth--see Professor Romanes on Panmixia, _Nature_, April 3, 1890.

[46] _Descent of Man_, p. 33.

[47] _Descent of Man_, p. 33.

[48] _Variation of Animals and Plants under Domestication_, i., 453.

[49] _Descent of Man_, p. 33.

[50] _Descent of Man_, p. 33.

[51] Wallace shows that the changes in our domestic animals, if spread over the thousands of years since the animals were first tamed, must be extremely insignificant in each generation, and he concludes that such infinitesimal effects of use and disuse would be swallowed up by the far greater effects of variation and selection (_Darwinism_, p. 436).

Professor Romanes has replied to him in the _Contemporary Review_ (August 1889), showing that this is no disproof of the existence of the minor factor, inasmuch as slight changes in each generation need not necessarily be matters of life and death to the individual, although their c.u.mulative development by use-inheritance might eventually become of much service. But selection would favour spontaneous variations of a similarly serviceable character. The slightest tendency to eliminate the extreme variations in either direction would proportionally modify the average in a breed. Use-inheritance appears to be so relatively weak a factor that probably neither proof nor disproof of its existence can ever be given, owing to the practical impossibility of disentangling its effects (if any) from the effects of admittedly far more powerful factors which often act in unsuspected ways. Thus wild ducklings, which can easily be reared by themselves, invariably "die off" if reared with tame ones (_Variation_, &c., i. 292, ii. 219). They cannot get their fair share in the compet.i.tion for food, and are completely eliminated.

Professor Romanes fully acknowledges that there is the "gravest possible doubt" as to the transmission of the effects of disuse (Letter on Panmixia, _Nature_, March 13, 1890).

[52] _Variation of Animals and Plants under Domestication_, ii. 287-289.

[53] _Descent of Man_, pp. 612, 131.

INHERITED INJURIES.

INHERITED MUTILATIONS.

The almost universal _non-inheritance_ of mutilations seems to me a far more valid argument _against_ a general law of modification-inheritance than the few doubtful or abnormal cases of such inheritance can furnish in its favour. No inherited effect has been produced by the docking of horses' tails for many generations, or by a well-known mutilation which has been practised by the Hebrew race from time immemorial. As lost or mutilated parts are reproduced in offspring independently of the existence of those parts in the parent, there is the less reason to suppose that the particular condition of parental parts transmits itself, or tends to transmit itself, to the offspring. So unsatisfactory is the argument derivable from inherited mutilations that Mr. Spencer does not mention them at all, and Darwin has to attribute them to a special cause which is independent of any general theory of use-inheritance.[54]

Darwin's most striking case--and to my mind the only case of any importance--is that of Brown-Sequard's epileptic guinea-pigs, which inherited the mutilated condition of parents who had gnawed off their own gangrenous toes when anaesthetic through the sciatic nerve having been divided.[55] Darwin also mentions a cow that lost a horn by accident, followed by suppuration, and subsequently produced three calves which had on the same side of the head, instead of a horn, a bony lump attached merely to the skin. Such cases may seem to prove that mutilation _a.s.sociated with morbid action_ is occasionally inherited or repeated with a prompt.i.tude and thoroughness that contrast most strikingly with the imperceptible nature of the immediate inheritance of the effects of use and disuse; but they by no means prove that mutilation in general is inheritable, and they are absolutely no proof whatever of a _normal_ and non-pathological tendency to the inheritance of acquired characters. Those who accept Darwin's special explanation of the supposed inheritance of mutilations, ought to notice that his explanation applies equally well under a theory which is strongly adverse to use-inheritance--namely, Galton's idea of the sterilization and complete "using up" of otherwise reproductive matter in the growth and maintenance of the personal structure.

Darwin's explanation of inherited mutilations--which, as he notes, occur "especially or perhaps exclusively" when the injury has been followed by disease[56]--is that all the representative gemmules which would develop or repair or reproduce the injured part are attracted to the diseased surface during the reparative process and are there destroyed by the morbid action.[57] Hence they cannot reproduce the part in offspring.

This explanation by no means implies that mutilation would _usually_ affect the offspring. On the contrary, in all ordinary cases of mutilation the purely atavistic elements or gemmules would be set free from any modifying influence of the non-existent or mutilated part. The gemmules--as in Galton's theory of heredity and with neuter insects--might be perfectly independent of pangenesis and the normal inheritance of acquired characters. Such self-multiplying gemmules without pangenesis would enable us to understand both the excessive weakness or non-existence of normal use-inheritance, and the excessive strength and abruptness of the effect of their partial destruction under special pathological conditions.

The series of epileptic phenomena that can be excited by tickling a certain part of the cheek and neck of the adult guinea-pig during the growth and rejoining of the ends of the severed nerve, are said to be repeated with striking accuracy of detail in the young who inherit mutilated toes; but as epilepsy is often due to some _one_ exciting cause or morbid condition, the single transmission of a highly morbid condition of the system might easily reproduce the whole chain of consequences and might also have caused the loss of toes.

The particulars of the guinea-pig cases are very inadequately recorded,[58] but the results are so anomalous[59] that Brown-Sequard's own conclusion is that the epilepsy and the inherited injuries are _not_ directly transmitted, but that "what is transmitted is the morbid state of the nervous system." He thinks that the missing toes may "possibly" be exceptions to this conclusion, "but the other facts only imply the transmission of a morbid state of the sympathetic or sciatic nerve or of a part of the medulla oblongata." Until we can tell what is transmitted, we are not in a position to determine whether there is any true inheritance or only an exaggerated simulation of it under peculiar circ.u.mstances. When the actual observers believe that the mutilations and epilepsy are not the cause of their own repet.i.tion, and when these observers guard themselves by such phrases as, "if any conclusion can at present be drawn from those facts," we who have only incomplete reports to guide us may well be excused if we preserve an even more p.r.o.nounced att.i.tude of caution and reserve.[60] The morbid state of the system may be wholly due to general injury of the germs rather than to specific inheritance.

Weismann suggests that the morbid condition of the nervous system may be due to some infection such as might arise from microbes, which find a home in the mutilated and disordered nervous system in the parent, and subsequently transmit themselves to the offspring through the reproductive elements, as the infections of various diseases appear to do--the muscardine silkworm disease in particular being known to be conveyed to offspring in this manner.

But whether we can discover the true explanation or not, inherited mutilations can hardly be accounted for as the result of a general tendency to inherit acquired modifications. How could a factor which seems to be totally inoperative in cases of ordinary mutilation, and only infinitesimally operative in transmitting the normal effects of use and disuse, suddenly become so powerful as to completely overthrow atavism, and its own tendency to transmit the non-mutilated type of one of the parents and of the non-mutilated type presented by the injured parent in earlier life? Does not so striking and abrupt an intensification of its usually insignificant power demand an explanation widely different from that which might account for the extremely slow and slight inheritance of the normal effects of use and disuse? Surely it would be better to suspend one's judgment as to the true explanation of highly exceptional and purely pathological cases rather than resort to an hypothesis that creates more difficulties than it solves.

THE MOTMOT'S TAIL.

The narrowing of the long central tail feathers of the motmot is attributed to the inherited effects of habitual mutilation (_Descent of Man_, pp. 384, 603). But in the specimens at South Kensington[61] the narrowness extends upwards much beyond the habitually denuded part, and the broadened end is the broadest part of the whole feather. If the inherited effect of an inch or two of denudation extends from three to six inches upwards, why has it not also extended two inches downwards so as to narrow the broadened end? The narrowness seems to be a mainly relative or negative effect produced by the broadening out of a long tapering feather at its end under the influence of s.e.xual selection.

Several other birds have similarly narrowed or spoon-shaped feathers and do not bite them. Is it not more feasible to suppose that this attractive peculiarity first suggested its artificial intensification, than to suppose that the bird began nibbling without any definite cause?

s.e.xual selection would then encourage the habit. Anyhow, it is as impossible to show that the mutilation preceded the narrowing as it is to show that tonsure preceded baldness.

OTHER INHERITED INJURIES MENTIONED BY DARWIN.

Darwin quotes some cases from Dr. Prosper Lucas's "long" but weak and unsatisfactory "list of inherited injuries."[62] But Lucas was somewhat credulous. One of his cases is that many girls were born in London without mammae through the injurious effect of certain corsets on the mothers. He also gives a long account of a Jew who could read through the thick covers of a book, and whose son inherited this "hyperaesthesia"

of the sense of sight in a still more remarkable degree (i. 113-119).

Evidently Lucas's cases cannot be accepted without some amount of reserve.

The cases of the three calves which inherited the one-horned condition of the cow, the two sons who inherited a father's crooked finger, and the two sons who were microphthalmic on the same side as their father had lost an eye, may be due to mere coincidence; or an inherited const.i.tutional tendency or liability might lead to somewhat similar results in parent and offspring[63]--just as the tendency to certain fatal diseases or to suicide may produce similar results in father and son, although the artificially-produced hanging or apoplexy obviously cannot be directly transmitted. That more than one of the offspring was affected does not render the chances against coincidence "almost infinitely great," as Darwin mistakenly supposes. It "frequently occurs"

that a man's sons or daughters may _all_ exhibit either a latent or a newly-developed congenital peculiarity previously unknown;[64] and the coincidence may merely be that one of the parents accidentally suffered a similar kind of injury--a kind of coincidence which must of course occasionally occur, and which may have been partly caused by a latent tendency. The chances against coincidence are indeed great, but the cases appear to be correspondingly rare.

Darwin acknowledges that many supposed instances of inherited mutilation may be due to coincidence; and there is apparently no more reason for attributing inherited scars, &c., to any special form of heredity than to the effect of the mother's imagination on the unborn babe--a popular but fallacious belief in corroboration of which far more alleged instances could be collected than of the inheritance of injuries.

As an instance of the coincidences that occur, I may mention that a friend of mine has a daughter who was born with a small hole in one ear, just as if it were already pierced for the earring which she has since worn in it. I suppose, however, that no one will venture to claim this as an instance of the inheritance of a mutilation practised by female ancestors, especially as such holes are not altogether unknown or inexplicable, though very rarely occurring low down in the lobe of the ear.[65]

Many cases are known of the inheritance of mutilations or malformations arising congenitally from some abrupt variation in the reproductive elements. In such cases as the one-eared rabbits, the two-legged pigs, the three-legged dogs, the one-horned stags, hornless bulls, earless rabbits, lop-eared rabbits, tailless dogs, &c., if the father or the mother or the embryo had suffered from some accident or disease which might plausibly have been a.s.signed as the cause of the original malformation, these transmitted defects would readily be cited as instances of the inheritance of an accidentally-produced modification.

The inheritance of exostoses on horses' legs may be the inheritance of a const.i.tutional tendency rather than of the effect of the parents' hard travelling. Horses congenitally liable to such formations would transmit the liability,[66] and this might readily be mistaken for inheritance of the results of the liability. An apparent increase in this liability might arise from greater attention being now paid to it, or from increased use of harder roads; or a real increase might be due to panmixia and some obscure forms of correlation.

QUASI-INHERITANCE.

Of course artificially-caused ill-health or weakness in parents will tend in a general way to injure the offspring. But deterioration thus caused is only a form of quasi-inheritance, as I should prefer to call it. Semi-starvation in a new-born babe is _not_ truly inherited from its half-starved mother, but is the direct result of insufficient nourishment. The general welfare of germs--as of parasites--is necessarily bound up with that of the organism which feeds and shelters them, but this is not heredity, and is quite irrelevant to the question whether particular modifications are transmitted or not.

Another form of quasi-inheritance is seen in the communication of certain infections to offspring. Not being transmitted by the action of the organism so much as in defiance of it, such diseases are not truly hereditary, though for convenience' sake they are usually so described.

A perversion or prevention of true inheritance is also seen in the action of alcohol, or excessive overwork, or any other cause which by originating morbid conditions in individuals may also injure the reproductive elements.

These forms of quasi-inheritance are, of course, highly important so far as the improvement of the race is concerned. So, too, is the fact that improved or deteriorated habits and thoughts are transmitted by personal teaching and influence and are c.u.mulative in their effect. But all this must not be confounded with the inheritance of acquired characters.

Cases of quasi-inheritance may perhaps be most readily distinguished from cases of true inheritance by the time test. When a modification acquired in adult life is promptly communicated to the child in early life or from birth, it may rightly be suspected that the inheritance, like that of money or t.i.tle, is not truly congenital, but is extraneous or even anti-congenital in its nature. Judged by such a standard, the inherited injuries in Brown-Sequard's guinea-pigs are only exceptional cases of quasi-inheritance, and are not necessarily indicative of any general rule affecting true inheritance.